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DISTRIBUTION AND ECOLOGY OF THE BROAD-TOOTHED RAT IN THE ACT RICHARD MILNER, DANSWELL STARRS, GREG HAYES AND MURRAY EVANS Conservation Research Technical Report 35 October 2016

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  • DISTRIBUTION AND ECOLOGY OF

    THE BROAD-TOOTHED RAT

    IN THE ACT

    RICHARD MILNER, DANSWELL STARRS,

    GREG HAYES AND MURRAY EVANS

    Conservation Research

    Technical Report 35

    October 2016

  • Technical Report 35

    Distribution and ecology of the

    Broad-toothed Rat in the ACT

    Richard Milner*, Danswell Starrs^, Greg Hayes* and Murray Evans*

    *Conservation Research, Environment and Planning Directorate,

    ACT Government, Canberra, ACT 2601

    ^Evolution, Ecology and Genetics, Research School of Biology,

    Australian National University, Canberra, ACT 0200

    Conservation Research

    Environment Division

    Environment and Planning Directorate

    October 2016

  • ISBN: 978-1-921117-50-0 Technical Report Series: ISSN 1320-1069: 35 © Australian Capital Territory, Canberra, 2016 Information contained in this publication may be copied or reproduced for study, research, information or educational purposes, subject to appropriate referencing of the source. This document should be cited as: Milner R, Starrs D, Hayes G and Evans M. 2016. Distribution and ecology of the broad-toothed rat in the ACT. Technical Report 35. Environment and Planning Directorate, ACT Government, Canberra. http://www.environment.act.gov.au Telephone: Access Canberra 13 22 81 Disclaimer The views and opinions expressed in this report are those of the authors and do not necessarily represent the views, opinions or policy of funding bodies or participating member agencies or organisations. Front cover: Broad-toothed Rat. G A Hoye © Australian Museum.

  • Distribution and ecology of the Broad-toothed Rat i

    Contents

    Summary ..................................................................................................................................................ii

    1 Introduction ..................................................................................................................................... 1

    2 Background ...................................................................................................................................... 2

    2.1 Description and Distribution ................................................................................................... 2

    2.2 Status ....................................................................................................................................... 2

    2.3 Habitat ..................................................................................................................................... 3

    2.4 Diet .......................................................................................................................................... 3

    2.5 Population dynamics ............................................................................................................... 3

    2.6 Breeding biology ...................................................................................................................... 4

    2.7 Threatening processes ............................................................................................................ 4

    3 Methods .......................................................................................................................................... 5

    3.1 Study sites ............................................................................................................................... 5

    3.2 Survey method ........................................................................................................................ 5

    3.3 Statistical Analyses .................................................................................................................. 6

    4 Results ............................................................................................................................................. 8

    4.1 Bogs ......................................................................................................................................... 8

    4.2 Distribution of M. fuscus ......................................................................................................... 8

    4.3 Habitat preference of M. fuscus .............................................................................................. 8

    4.4 Feral animal disturbance ......................................................................................................... 9

    4.5 Observer bias ........................................................................................................................... 9

    5 Discussion ...................................................................................................................................... 18

    5.1 Distribution, status and abundance ...................................................................................... 18

    5.2 ACT Habitat ............................................................................................................................ 18

    5.3 Bog size .................................................................................................................................. 19

    5.4 Vegetation composition ........................................................................................................ 19

    5.5 Threats and Management recommendations ...................................................................... 19

    5.6 Conclusion ............................................................................................................................. 22

    Acknowledgements ............................................................................................................................... 22

    References ............................................................................................................................................. 22

    Appendix................................................................................................................................................ 25

  • ii Distribution and ecology of the Broad-toothed Rat

    Summary

    The Broad-toothed Rat Mastacomys fuscus is a small to medium sized rodent of the family Muridae.

    Within the ACT the species is thought to occur in subalpine swamps and the associated riparian

    heathland, sedgeland and grassland environments above 1400 m. M. fuscus is recognised nationally

    as a declining species endemic to alpine and sub-alpine regions of south-eastern Australia, including

    Tasmania, Victoria, NSW and the ACT. The species is identified as being at extreme risk to global

    warming due to shifts in composition and distribution of alpine vegetation communities. Climate

    change has been identified as a key threatening process to the long-term viability of the species. In

    NSW M. fuscus is classified as vulnerable, with one of the two populations, Barrington Tops, classified

    as endangered. In Victoria the species is classified as near threatened and in the ACT, where only

    limited information is available on the species, M. fuscus has no special protection status.

    The purpose of this study was to determine the presence or absence of the M. Fuscus in known

    habitat for the species (sphagnum moss bogs and fens) in the ACT, and to identify key ecological

    attributes that determine the presence of the species. A key aim of this study was to establish a long

    term monitoring program to assess long-term trends in the species’ abundance and distribution,

    which may be affected under projected climate change scenarios.

    Fourteen ACT alpine bogs were surveyed across Namadgi National Park, which covers the southern

    half of the ACT and adjoins Kosciusko National Park. Scat searches were undertaken in 27-132

    quadrats per bog to determine the species presence/absence, relative abundance and habitat

    preference. For each quadrat, information on M. fuscus presence/absence, vegetation type, feral

    animal disturbance and distance to drainage lines was recorded. Bog size and distance to the nearest

    occupied bog was also recorded.

    Results indicate M. fuscus presence across 13 of the 14 sites: Cotter Gap, Cheyenne, Rocks Flat,

    Snowy Flat, Ginini West, Big Creamy, Ginini East, Tom Gregory, Little Bimberi, Rotten Swamp, Murray

    Flat, Leura and Top Flat. The species showed a positive habitat preference for vegetation

    communities dominated by heath, sedge and Poa, and was also identified in a further nine dominant

    vegetation types, including Eucalyptus pauciflora (snowgum) woodland. Vegetation composition

    varied with bog size and M. fuscus detection rate increased with increasing bog size. Distance from

    drainage lines was also important, with detection rates increasing with proximity to drainage lines.

    Finally the presence of feral animals such as foxes, rabbits and pigs significantly influenced detection

    rates, with M. fuscus detection rate decreasing as signs of feral animal disturbance increased.

    This study provides land managers with a baseline assessment of M. fuscus condition in the ACT

    against which future changes can be assessed and key-threatening processes can be monitored.

    The results of this study have also been published in:

    Milner R, Starrs D, Hayes G and Evans M. 2014. Distribution and habitat preference of the broad-

    toothed rat (Mastacomys fuscus) in the Australian Capital Territory, Australia. Australian

    Mammalogy. http://dx.doi.org/10.1071/AM14031

  • Distribution and ecology of the Broad-toothed Rat 1

    1 Introduction

    The Broad-toothed Rat, Mastacomys fuscus, is a cryptic native rodent that inhabits montane,

    subalpine and alpine areas, usually in close association with water bodies. In the ACT M. fuscus is

    known to occur in sphagnum moss bogs, although surveys for the species have not been conducted

    for over two decades and only limited information exists on its distribution and abundance. The

    species is known to have declined in other jurisdictions, and is listed as threatened in NSW and

    Victoria. Climate change has been identified as a key threatening process to the long-term viability of

    the M. fuscus due to shifts in composition and distribution of alpine vegetation communities.

    The broad goal of this study was to gain a better understanding of the distribution, abundance,

    ecology and conservation status of M. fuscus in the ACT to support appropriate management

    decisions for the species and its habitat.

    Specific aims of study were to:

    Determine the distribution, abundance and status of the Broad-toothed Rat Mastacomys

    fuscus in the ACT.

    Identify key ecological attributes that determine or influence the likelihood of the species

    occurrence.

    Establish a baseline and method for long term monitoring to assess the effects of climate

    change on the abundance and distribution of the species.

  • 2 Distribution and ecology of the Broad-toothed Rat

    2 Background

    2.1 Description and Distribution

    The Broad-toothed Rat, Mastacomys fuscus, is a small to medium sized rodent of the family Muridae

    (Happold 2002). Adult head-body length ranges from 142–175 mm, tail length ranges from 100–130

    mm and adult weight ranges from 95–145 g. The species is found across a range of altitudes from sea

    level to 2200 m where habitat is suitable in the Dandenong and Otway Ranges, Victorian Alps, Snowy

    Mountains, Barrington Tops and Tasmania;. Across Snowy Mountains and Barrington Tops

    populations, the species habitat is characterised by mean annual temperatures 1000 mm (Green and Osborne 2003). While the species is currently restricted to

    patches of apparently optimal habitat, fossil records indicate a more extensive distribution during the

    Pleistocene when the climate was cooler (Ride 1956).

    Within the ACT there is limited information on the distribution, abundance and ecology of M. fuscus.

    Only three surveys in the ACT have successfully detected the species, all of these were undertaken

    prior to 1990. In 1973 Eberhard and Schulz (1973) captured a single M. fuscus at Murrays Gap when

    undertaking a vertebrate survey of the Cotter River catchment. In 1986, Lintermans (1986)

    undertook a survey of M. fuscus across six ACT sites (Murrays Gap, Cheyenne Flats, Stockyard Saddle,

    saddle below Mt Gingera, Mt Gingera and Snowy Flats) from which only one individual was captured

    (at Murray Gap). From 1988 to 1989, Green and Osborne (2003) surveyed 25 areas within or

    bordering the ACT with M. fuscus recorded within 12 of the sites: Hanging Flat, Murrays Gap, Mt

    Murray, Ginini Flats, Cheyenne Flats, Stockyard Flats, Brumby Flats, 80 Acres, Blackfellows Gaps,

    Rolling Ground Gap, Mt Bimberi and Mt Scabby. Runways and scats of M. fuscus have been observed

    at Ginini Flats, Cheyenne Flats, Morass Flats and Snowy Flats (M Evans pers. obs.; D Whitfield pers.

    obs.).

    2.2 Status

    M. fuscus is recognised nationally as a declining species endemic to alpine and sub-alpine regions of

    south-eastern Australia, including Tasmania, Victoria, NSW and the ACT. In Victoria M. fuscus is listed

    as Near Threatened, and in NSW the species is listed as Threatened (Vulnerable) though one of the

    populations (Barrington Tops) is classified as Endangered. M. fuscus is not listed as threatened in the

    ACT, where there is a paucity of information on the species.

    M. fuscus is identified as being at extreme risk to global warming due to loss of habitat (Sphagnum

    moss bogs and other wet seepage areas) resulting from shifts in the composition and distribution of

    alpine vegetation communities (DECC NSW 2007), and accordingly climate change has been

    identified as a key threatening process to the long-term viability of M. fuscus. Brereton and

    colleagues (1995) predicted that with a 1oC rise in temperature the range of M. fuscus would

    decrease by 36% and with a 3 oC rise in temperature it would decrease by 75%. In 2005, O’Brien

    showed that extinction of M. fuscus at Barrington Tops is expected under almost any climatic

    warming scenario.

  • Distribution and ecology of the Broad-toothed Rat 3

    2.3 Habitat

    M. fuscus is associated with a variety of habitat types, including rocky outcrops, tall tussock

    grasslands, reeds, sedges and low spreading heath shrubs, usually within 15 m of watercourses (K

    Green pers. comm.). The species requires dense ground cover and an abundance of green grass,

    including snow grasses (Poa spp). In the ACT, M. fuscus is apparently associated with subalpine

    swamps and associated riparian heathland, sedgeland and grassland environments above 1000 m,

    with a mean annual rainfall greater than 1000 mm and average temperature

  • 4 Distribution and ecology of the Broad-toothed Rat

    winter (Bubela et al. 1991). Female home range size is determined by metabolic requirements such

    as food quality and food density (Mace et al. 1983), while male home range size is also subject to

    sexual selection for increased access to females, with home range decreasing as female reproductive

    receptiveness decreases (Bubela et al. 1991). While M. fuscus do not appear to display territorial

    defence, there is evidence that both sexes occupy an exclusive/solitary space that averages 802m2

    within their home range during January to autumn (Bubela et al. 1991).

    2.6 Breeding biology

    M. fuscus display female-only parental care and are believed to be either polygamous or

    promiscuous (Happold 2011). Mating occurs in mid-late October and is trigged by a gradual increase

    in ambient temperature, snowmelt, decreased frequency of frost, increasing day length and the

    growth of new grass (Bubela and Happold 1993). Breeding nests are built under dense shrubs,

    mattered stems, boulders and fallen logs and are similar to overwinter nests – they have a single

    entrance and a single internal cavity up to 30 cm in diameter (Happold 2011). Gestation period is 35

    days (Calaby and Wimbush 1964) and all females synchronously give birth to their first litter in the

    last week of November or first week of December (Happold 1989). Births of the second and

    sometimes third litters are spread out over several weeks from December to early March and litter

    size ranges from 1–4 (Bubela et al. 1991; 1993). Young M. fuscus start to eat grass at three weeks and

    are weaned at five weeks (Happold 2011) and sexual maturity is reached at 8–10 months.

    2.7 Threatening processes

    Key threatening process recognised by the NSW Threatened Species Conservation ACT 1995 that are

    relevant to M. fuscus include:

    predation by European Red Fox Vulpes vulpes and the Feral Cat Felis catus;

    competition and grazing by feral European Rabbits Oryctalagus cuniculus;

    competition, disease transmission and habitat degradation by Feral Pigs (Sus scrofa);

    invasion of exotic perennial grasses;

    climate change resulting in loss of sub-alpine and alpine habitat, and;

    spread of the plant root fungus Phytophthora cinnamomi.

  • Distribution and ecology of the Broad-toothed Rat 5

    3 Methods

    3.1 Study sites

    Potential habitat for M. fuscus was identified remotely using aerial photographs, vegetation maps

    (i.e. sphagnum shrub bog, sod tussock Poa grassland, Poa-Danthonia grassland, and Restiad bog

    ArcGIS layers – CPR GIS system), ACT Wildlife Atlas records, and expert knowledge (D Whitfield pers.

    comm.). Fourteen study sites were identified, including: Cotter Source, Top Flat, Rotten Swamp, Big

    Creamy, Rocks Flat, Cotter Gap, Tom Gregory, Murray Gap, Little Bimberi, Leura, Snowy Flat,

    Cheyenne Flat, Ginini West and Ginini East.

    3.2 Survey method

    Surveys were undertaken in early autumn (March) 2013. Area-based searches for scats (faecal

    pellets) were undertaken in 27–130 circular quadrats (2 m radius) per study site. Quadrats were

    positioned 20 m apart along 5–12 straight-line transects. Line transects were positioned at right

    angles to the streams and elevational gradient (i.e. transects cut across contours; Map 1) and were

    placed 50 m apart for study sites/bogs > 35,000 m2 and 25 m apart for study sites/bogs

  • 6 Distribution and ecology of the Broad-toothed Rat

    3.3 Statistical Analyses

    The proportion of quadrats with M. fuscus scats was plotted against bog size (km2). Linear regression

    was used to determine if a significant relationship existed between M. fuscus abundance and bog

    size. Principle component analysis on the covariance matrix was used to explore variation in

    vegetation types between bogs. Proportional cover of vegetation across 11 bogs was explored (three

    bogs were excluded due differences in the method of reporting of vegetation types, which did not

    enable comparison of vegetation cover data). Principle components 1 and 2 were plotted, and

    distances across Euclidean space interpreted from raw data. Bog size and principle components were

    plotted to examine if principle components corresponded to bog size. Average proportional cover (±

    SE) of each vegetation type was plotted to examine the relative importance of each vegetation type.

    A habitat selectivity analysis (Manly et al. 2002) was used to examine M. fuscus preference for

    specific habitat types. We assumed that vegetation preferences would not vary between bogs, so

    data across all bogs was combined to improve sample sizes for each vegetation category. A Type I

    (Manly et al. 2002) selectivity analysis was conducted in the R package AdehabitatHS (Calenge 2013).

    The proportion of available habitat was calculated as the proportion of quadrats containing each

    vegetation type, and M. fuscus usage was estimated as the count of quadrats that contained M.

    fuscus scats corresponding to each vegetation type. Habitat selection indices (Wi) were calculated for

    each vegetation type. Overall significance of habitat selection was determined by the log-likelihood

    statistic (Manly et al. 2002).

    A total of three additional observers assisted the lead author in identifying M. fuscus scats at five

    bogs (Cheyenne, Ginini East, Leura, Snowy Flat and Tom Gregory). Only one additional observer was

    present at any one time. Observer bias was explored by examining the relative proportion of M.

    fuscus scats discovered by an observer, after controlling for bog-level differences in M. fuscus

    abundance. If observer bias was present, it would be expected to manifest itself through differences

    in M. fuscus scat detection between observers. M. fuscus scat abundance was modelled against

    observer, with bogs included as a random factor to account for bog-level differences in M. fuscus

    abundance.

    The relationship between proportion of quadrats with M. fuscus scats and distance (transformed by

    the square-root to meet assumptions of statistical normality) from nearest drainage line was

    explored in a GLMM framework. Bog ID was included as a random factor, to account for possible

    differences between bogs. In a similar manner, the relationship between M. fuscus abundance and

    incidence of feral animal disturbance was explored in a mixed statistical model. All feral animal

    disturbance indicators (fox, rabbit and pig) were combined into a single category. While it is plausible

    that the relationship between M. fuscus and these species differs due to ecological relationships

    (competition, predation), we chose to combine these categories due to limited data for each feral

    species. Mixed effects models were implemented in R with the package lme4 (Bates et al. 2011).

    Plots were produced in R, with the fixed factors plotted for each level of the random factor (bog ID).

    A final model, fitting M. fuscus abundance against the three positively selected vegetation types,

    feral animal disturbance and square-root of linear distance to drainage (m). Bog ID was included as a

    random factor. The presence of spatial autocorrelation in the model residuals was explored by way

    of Moran’s I, in the R package ‘ape’ (version 3.2, Paradis et al. 2003). No significant spatial

    autocorrelation was detected (P = 0.84). Model fitting was performed in R (version 2.13; R

    Development Core Team, 2011). Model selection was conducted in ‘glmulti’ (Calcagno and de

  • Distribution and ecology of the Broad-toothed Rat 7

    Mazancourt 2010), using an exhaustive screening of all possible models with 2-way interactions. The

    best model was identified using AIC. Figures were produced in Sigmaplot (version 10; Systat Software

    Inc., London).

    Map 1. Survey design and site establishment for Mastacomys fuscus in the ACT (Snowy Flat).

  • 8 Distribution and ecology of the Broad-toothed Rat

    4 Results

    4.1 Bogs

    Fourteen alpine bogs in Namadgi National Park were surveyed for M. fuscus scats and vegetation

    characteristics in March, 2013. A total of 13 vegetation/habitat types were recorded from quadrats

    across all bogs. A Principle Components Analysis revealed that the proportion of vegetation type

    could provide separation between bogs in Euclidean space, with PC1 and PC2 explaining 46.39% and

    20.05% of the total variation in vegetation between bogs respectively (Figure 1). Proportion of

    vegetation in the categories of Exotic plants, Empodisma, HeathL, Sphagnum, Richea, and HeathT

    were particularly informative in describing differences between bogs, whereas the proportion for

    categories of Poa, rocks, Forbs and woodland were not (Figure 1a). Tom Gregory and Cotter Gap bogs

    differed from the other bogs due to their relatively high presence of exotic species (33% and 16.7%,

    respectively), while Little Bimberi, Cheyenne Flat and Ginini differed from the other bogs in their

    relatively higher proportions of Sphagnum and Richea (Figure 1b). Poa was the most abundant

    vegetation type in 11 of the 14 bogs, being present in 75% ± 3.63 SE of all quadrats (Figure 2).

    HeathL, HeathM, Empodisma and Sedge were present in more than 20% of quadrats on average,

    across all bogs (Figure 2). Conversely, the categories of Forbs, Richea and Juncus, and rocks were

    relatively rare in bogs, each being present at less than 5% of all quadrats. The abundance of exotic

    plant species was relatively low in bogs; this category was recorded at just three bogs (Tom Gregory,

    Cotter Gap and Murray Gap), with bog Tom Gregory having the highest proportion of quadrats

    containing exotic plants (33%).

    4.2 Distribution of M. fuscus

    Across all bogs, M. fuscus scats were found in 26.5% of quadrats examined. At the landscape-scale,

    M. fuscus density (proportion of quadrats within a bog that had M. fuscus scats) varied greatly

    between bogs (Figure 3a). M. fuscus density was greatest at Cotter Gap bog (69%), and none were

    present at Cotter Source bog (0%; Table 1; Figure 3a). The probability of detecting M. fuscus scats

    was not related to bog size, however when excluding an outlier (bog Cotter Gap), which supported

    an unusually high density of M. fuscus, there was a significant positive relationship between bog size

    and M. fuscus density (Figure 3b). PC1 was negatively correlated to bog size, suggesting that

    vegetation types vary by bog size (Figure 4). However, M. fuscus abundance was not related to PC1

    or PC2.

    4.3 Habitat preference of M. fuscus

    Pooling data across all bogs, habitat selectivity indices (Wi) suggested that M. fuscus show

    preferences for specific vegetation types (log-likelihood statistic: X2 = 31.75, df = 12, P < 0.001)

    (Figure 5). Specifically, after bonferroni correction for multiple comparisons, M. fuscus scat presence

    was found to be significantly positively associated with medium height heath (HeathM) (Figure 5;

    Appendix Figure C). M. fuscus also showed weak but non-significant preferences for Poa and Sedge

    (Appendix Figure D-E). Conversely, there was a weak, non-significant negative association with

    Empodisma, woodland and low heath (HeathL) (Appendix Figure F-G). Apparent preferences for

    rarely occurring vegetation types (such as Forbs, Richea, Juncus) and rock, were given a low

    confidence due to the low abundance of these vegetation types and hence small sample sizes. The

  • Distribution and ecology of the Broad-toothed Rat 9

    probability of M. fuscus scats being observed was negatively related to straight-line distance from

    drainage lines, with the likelihood of detecting M. fuscus scats declining by 1.1% per m distance from

    a drainage line (Figure 6; Table 2). This relationship was found to be reasonably consistent between

    bogs, after accounting for the high bog-level variance (59%) (Table 2; Figure 6).

    4.4 Feral animal disturbance

    The probability of detecting M. fuscus scats decreased significantly if evidence of feral animal

    disturbance was present (Table 3). Feral animal disturbance was observed in 11.2% of quadrats

    across all bogs, with Cotter Source having the highest incidence of feral animal disturbance (31.1%;

    Table 1). Of the observed feral animal disturbance in bogs, evidence of pigs was most common

    (80.2%), with fox and rabbit representing 18.1% and 1.7% respectively. With one exception,

    modelling revealed a consistent decrease in M. fuscus density at the bog level, suggesting that feral

    animals have a consistent, negative effect on M. fuscus abundance. The exception occurred at Cotter

    Gap, where M. fuscus density increased with feral animal disturbance (Figure 7).

    A model explaining M. fuscus scat presence with respect to the three most preferred vegetation

    types, distance from drainage and presence of feral animal disturbance was found to provide the

    best fit, compared to models with a subset of these variables (Table 4). Likelihood of detecting M.

    fuscus scats declined with distance from drainage line, and presence of feral animal disturbance.

    Conversely, likelihood of detection increased significantly with the presence of Poa, Sedge and

    medium height heath (Table 4). All interaction terms were found to be non-significant when the main

    effect was present within the model. The random factor (bog ID) explained 34% of variation.

    4.5 Observer bias

    Observer bias was found to be non-significant. After accounting for bog-level variance, there were no

    significant differences in scat detection rates among observers (Table 5). This is evidenced by the

    minor differences between observer coefficients, relative to standard errors (Table 5).

  • 10 Distribution and ecology of the Broad-toothed Rat

    Figure 1. Principle component analysis of 13 vegetation types recorded across 11 alpine bogs. a) Ordination plot of principle component 1 and principle component 2, explaining 46.39% and 20.05% of variation in vegetation types, respectively, and b) vector plot, showing the relationships between bogs based upon similarities and differences in vegetation types.

  • Distribution and ecology of the Broad-toothed Rat 11

    Figure 2. Average percentage of each vegetation type present at each site along transects for 11 alpine bogs. Poa was the most common vegetation type, followed by Empodisma and medium heath.

    Table 1. Summary of survey effort, M.fuscus detection rate and feral animal disturbance per bog.

    Bog Transects Plots Plots occupied Detection rate %

    Feral animal disturbance %

    Cotter Gap 10 42 29 69 17

    Cheyenne 11 77 33 43 4

    Rock Flat 8 70 24 34 13

    Snowy Flat 11 133 44 33 6

    Ginni West 12 100 30 30 13

    Big Creamy 10 84 21 25 10

    Ginni East 8 131 31 24 18

    Tom Gregory 11 78 18 23 5

    Rotten Swamp 6 72 13 18 8

    Little Bimberi 11 61 11 18 11

    Murray Gap 10 63 10 16 2

    Leura 5 28 4 14 7

    Top Flat 10 53 7 13 19

    Cotter Source 10 45 0 0 31

  • 12 Distribution and ecology of the Broad-toothed Rat

    Figure 3. a) Proportion of spots that contained M. fuscus scats, by bog ID, and b) relationship between index of M. fuscus abundance (from panel a) and bog size (km2). Linear model is fitted to 13 bogs with an outlier (Cg; Cotter Gap) excluded (open circle).

  • Distribution and ecology of the Broad-toothed Rat 13

    Figure 4. Bog size (km2) plotted against principle component 1 (PC1) from the PCA analysis. An outlier (bottom left; bog Lb; Little Bimberi) was excluded from the linear model. When included, model fit is still significant (F1,9 = 9.37, P = 0.01), although quality of fit is reduced (R2 = 0.51).

    Figure 5. Habitat selection index (Wi) for M. fuscus, averaged across 11 alpine bogs. Vegetation types greater than the dashed line show positive selection by M. fuscus, while those below the dashed line are negatively selected by M. fuscus.

  • 14 Distribution and ecology of the Broad-toothed Rat

    Table 2. Summary of general linear mixed effects model on likelihood of detecting M. fuscus scats, and square-root linear distance from drainage, across 14 alpine bogs in Namadgi NP. Bog ID was included as a random factor (random intercept and slope) to account for bog-level variance.

    Fixed Factors Coefficient SE Z P

    Intercept -0.66 0.245 -2.698 0.007

    DistDrainL1/2 -0.088 0.026 -3.38

  • Distribution and ecology of the Broad-toothed Rat 15

    Table 3. Summary of general linear mixed effects model of likelihood of detecting M. fusucs scats and incidence of feral animal disturbance, across 14 alpine bogs in Namadgi NP. Bog ID was included as a random factor (random intercept and slope) to account for bog-level variance.

    Fixed Factors Coefficient SE Z P

    Intercept -1.07 0.22 -4.9

  • 16 Distribution and ecology of the Broad-toothed Rat

    Table 4. Summary of general linear mixed effect model on likelihood of detecting M. fuscus scats and key vegetation types (Poa, heathM and sedge), distance from drainage, and incidence of feral animal disturbance, across 11 alpine bogs in Namadgi NP. Bog ID was included as a random factor (random intercept) to account for bog-level differences in mean M. fuscus abundance.

    Fixed Factors Coefficient SE Z P

    Intercept -1.24 0.3 -4.08

  • Distribution and ecology of the Broad-toothed Rat 17

    Map 2. ACT Mastacomys fuscus distribution 2013.

  • 18 Distribution and ecology of the Broad-toothed Rat

    5 Discussion

    5.1 Distribution, status and abundance

    Mastacomys fuscus was detected across 13 of 14 bogs surveyed in the ACT. Detection rate increased

    as bog size increased and decreased as distance to drainage line increased. The species showed a

    positive habitat preference for heath, sedge and Poa dominated vegetation types. Feral animal

    disturbance negatively influenced detection rate in almost all bogs.

    The 14 ACT subalpine bogs surveyed in this study ranged in altitude from 1028 m to 1629 m, and

    incuded Cotter Gap, Cheyenne, Rocks Flat, Snowy Flat, Ginini West, Big Creamy, Ginini East, Tom

    Gregory, Little Bimberi, Rotten Swamp, Murray Flat, Leura and Top Flat. M. fuscus was detected in 13

    of these bogs; the only bog surveyed that did not support detectable levels of M. fuscus was the

    Cotter Source bog. Feral animal disturbance in this bog was high with over 31% of quadrats showing

    some sign of feral animal disturbance. The results of this study increase the number of known sites to

    have supported M. fuscus in the ACT from 14 to 23 (Map 2), thereby extending the species’ known

    distribution in the region. Previous evidence of the species in the ACT comes from only three surveys

    from over 23 years ago and opportunistic observations at some sites of runways and scats. Due to

    the nature of the previous surveys (i.e. presence/absence) it is not possible to identify trends in

    population size over the past 40 years. The information collected in the current study can be used as

    a baseline for longer-term monitoring of M. fuscus distribution, abundance and habitat use and also

    for detecting shifts in the composition and distribution of bog vegetation communities, which might

    be expected in response to climate change. Shifts in composition and distribution of alpine

    vegetation communities due to climate change have been identified as a key threat to the

    persistence of M. fuscus across its range (DECC NSW 2007).

    5.2 ACT Habitat

    M. fuscus was found to occur across a variety of habitat types in the ACT including: rocky outcrops,

    tussock grasslands, sedgelands and heathlands, frequently within close proximity to watercourses.

    Evidence of woodland habitat use in areas adjacent to subalpine bogs and grasslands was also found,

    although results of a habitat selectivity analysis indicated a weak negative association with the

    vegetation type, most likely due to a lack of appropriate ground cover. Overall, M. fuscus was

    strongly associated with sites dominated by heath (50–150 cm high) and/or areas dominated by Poa

    and sedges. Surprisingly M. fuscus showed a weak, negative association with low spreading heath,

    which would be expected to provide excellent cover and protection from predators (DECC NSW

    2007). However, the accuracy of this result is uncertain because scat detectability may have been

    reduced due to poor visibility.

    Proximity to drainage lines is apparently important to M. fuscus, with the likelihood of detecting the

    species decreasing 1.1% every metre away from a drainage line. These results are consistent with

    observations from Kosciuzsko National Park that indicate the species is most frequently encountered

    within 15 m of watercourses (K Green pers. comm.). Drainage lines are likely to support a higher

    density and complexity of vegetation, as well as a higher abundance of grasses, both of which are key

    habitat attributes for the species. M. fuscus feed primarily on monocotyledons and is generally

    associated with dense vegetation cover (Carron et al. 1990). Nil spatial autocorrelation in model

  • Distribution and ecology of the Broad-toothed Rat 19

    residuals indicate that in the present study, linear distance from nearest drainage is an adequate

    predictor of M. fuscus distribution in relation to drainages.

    5.3 Bog size

    The detection rate of M. fuscus increased as bog size increased. In a number of animal species,

    habitat patch size has been shown to influence population persistence (Banks et al. 2007). Larger

    habitat patches are likely to support larger populations as well as provide a greater ‘target’ for

    individuals dispersing into the area (Hanski and Simberloff 1997). Interestingly O’Brien et al (2008)

    showed that there is no evidence of patch size influencing population persistence, although habitat

    patch sizes for M. fuscus in the Barrington Plateau tended to be larger than those found in the ACT.

    There is evidence of populations of M. fuscus in the ACT region declining in isolated swamps, most

    likely the result of reduced dispersal success through predation by introduced predators (O’Brien et

    al. 2008). M. fuscus are good dispersers and have been shown to recolonise high quality patches

    following local extinctions, though long distance movements are uncommon, especially if riparian

    cover or creek lines are absent (Bubela et al. 1991; Happold 1995; O’Brien 2008). Drainage systems

    between bogs should therefore be managed as a key habitat resource for the species, aiding

    dispersal and thereby the persistence of the species in the region.

    5.4 Vegetation composition

    Vegetation composition varied with bog size. Across all bogs surveyed, Poa grass was the most

    abundant species recorded, followed by heath (

  • 20 Distribution and ecology of the Broad-toothed Rat

    association with both Poa grasslands and sedgelands and negative association with sphagnum.

    However, with elevated summer temperatures, woody vegetation is likely to increase in alpine areas

    (Williams and Costin 1994). M. fuscus were shown in this study to utilise woodlands adjacent to high

    quality habitat, though results of the habitat selectivity analysis showed that M. fuscus tend to avoid

    woodland habitat and therefore the expansion of woody vegetation may reduce available optimal

    habitat and lead to an overall reduction in habitat complexity and structure. Furthermore, while

    restricted altitudinal movement is recognised as a significant impact of climate change for M. fuscus

    populations in the Barrington Plateau (NSW), the significance of such processes in ACT populations

    are currently unknown. Populations of M. fuscus have been identified across a range of altitudinal

    gradients (1028 m to 1629 m) and, unlike the Barrington Plateau population, altitudinal movement

    for the species in the ACT is less restricted. Indeed, populations of M. fuscus have been identified at

    relatively low altitudes in the ACT (e.g. 1028 m).

    Fire

    An increase in fire frequency within alpine bog communities is predicted to increase the number of

    peaty sedgelands, grasslands and heath, while decrease the number of Sphagnum peatlands

    (MacDonald 2009). While an increase in sedgeland, grassland and heath vegetation is likely to

    increase available habitat for M. fuscus in the long-term, high intensity fire is likely to significantly

    impact populations of M. fuscus in periods immediately following fires through direct mortality and

    loss of resources (e.g. food and shelter). Even in unburnt areas, predation by foxes on M. fuscus and

    other small mammals has been shown to increase following fires, with predation concentrated in

    areas still carrying populations of small mammals (Green and Sanecki 2006). The wildfire in 2003

    burnt most of the habitat for M. fuscus, though individuals managed to persist in unburnt habitat

    patches as evidenced by recent scats and runways in these patches (Carey et al. 2003). While M.

    fuscus are good dispersers and have been shown to recolonise high quality patches following

    extinction (i.e. following high intensity fires), re-colonisation following such events will depend on the

    level of connectivity between populations as well as proximity from source populations. While the

    Top Flat population is relatively isolated from other known populations of M. fuscus, the population

    is well connected to other populations via suitable dispersal habitat along the Cotter River.

    Nevertheless, actions to reduce high intensity fire risk should be incorporated into the management

    of such relatively isolated sites.

    It should be noted that a number of heath species associated with M. fuscus require fire for

    reproduction. A low intensity fire frequency of 10–40 years is likely to be appropriate for maintaining

    or improving heath habitat (NPWS 2005) for M. fuscus. Sphagnum peatlands (an Endangered

    Ecological Community under the EPBC Act 1999) and surrounding woodland habitat should not be

    subject to this fire regime, and instead fire should be excluded from these habitats, particularly the

    fire-sensitive Sphagnum moss bog communities. While peatlands in good condition are relatively

    resistant to low intensity fire, increased evaporation and reduced rainfall with a changing climate

    (Lucas et al. 2007) is predicted to dry out Sphagnum peatlands and significantly increase the risk and

    impact of fire within the community (Macdonald 2009).

    Habitat degradation through feral animal disturbance

    Habitat degradation by feral pigs and rabbits poses a significant threat to populations of M. fuscus in

    the ACT. Damage caused by rabbits and pigs was recorded in all 14 bogs surveyed and was

    significantly correlated with lower M. fuscus detection rate across all bogs except Cotter Gap. Feral

  • Distribution and ecology of the Broad-toothed Rat 21

    pig and rabbit activity is likely to result in erosion, a reduction in food resources and a change in

    habitat structure and composition (Green and Osborne 2003).

    Feral animal activity is likely to contribute significantly to the establishment and spread of weeds and

    it is possible that feral pigs could play a key role in the spread of Phytophthora. Phytophthora poses a

    significant threat to the species habitat – leading to a loss of heath and other vegetation essential to

    the species survival. Bogs with M. fuscus and other threatened species should be priority target areas

    for landscape scale feral animal control programs in Namadgi National Park.

    Weeds

    The invasion of exotic plant species into key M. fuscus habitat poses a significant threat to the

    species survival. The invasion of exotic grasses is likely to result in the loss of important ground layer

    vegetation such as Snow grass with the potential to reduce the availability of that food source for M.

    fuscus. Yorkshire fog Holcus lanatus was recorded at relatively high density at the Cotter Gap bog,

    though there is currently no evidence to indicate that this introduced species is having an adverse

    affect on the population as M. fuscus detection rate in this bog was high. H. lanatus has, however,

    been identified as a potential threat to populations of M. fuscus in the Barrington Tops where there

    is concern that this grass may displace significant native ground layer species such as Snow Grass,

    which M. fuscus is known to rely heavily on as a food resource (DECC NSW 2007). Careful monitoring

    of the Cotter Gap bog, and other bogs that can be accessed by the public on foot or by vehicle, is

    recommended to ensure that infestations of H. lanatus and other weeds do not increase and that the

    presence of the species is not having an adverse affect on the population of M. fuscus. It should be

    noted that H. lanatus is unlikely to be a food resource for M. fuscus.

    Competition

    A number of both exotic and native species compete with M. fuscus for food, shelter and space.

    Evidence of rabbits was recorded in nine of the 14 sites surveyed, with disturbance highest at Cotter

    Gap, Cotter Source and Tom Gregory. Rabbits compete directly with M. fuscus for food; and grazing

    by rabbits can result in the removal or modification of tussock structure that provides shelter and

    nesting habitat for M. fuscus. Control of rabbits and foxes is likely to be most effective when

    undertaken concurrently at a landscape scale, and control of these species is particularly important

    at the following three bogs: Cotter Source, Cotter Gap and Tom Gregory.

    Competition for resources from native species, such as the Bush Rat Rattus fuscipes, may pose an

    additional competitive threat to M. fuscus. For example, a significant increase in the number of R.

    fuscipes trapped in Nolans Swamp (NSW) coincided with a dramatic decrease in the number of M.

    fuscus captured, indicating a potential dominance of R. fuscipes over M. fuscus (Keating 2003). R.

    fuscipes were detected in four of the 14 bogs surveyed (incl. Cheyenne, Murray Gap and Ginini East

    and West) in the ACT. However, R. fuscipes is naturally sympatric with M. fuscus and therefore it is

    unknown if and under what circumstances R. fuscipes poses a potential competitive threat to M.

    fuscus in ACT populations.

    Predation

    M. fuscus fall within the Critical Weight Range (CWR; Burbidge and McKenzie 1989) and the

    Vulnerable Terrestrial Vertebrate category (Smith and Quin 1996) indicating a high risk of population

    decline and extinction through predation from foxes and cats. There is also evidence of selective

  • 22 Distribution and ecology of the Broad-toothed Rat

    predation by foxes on M. fuscus over other sympatric CWR rodent species. For example, Green

    (2002) showed that M. fuscus in the alpine regions of the Snowy Mountains were five times more

    frequently found in fox scats than were R. fuscipes despite occurring at relatively similar densities.

    The disparity in predation rate has been attributed to behavioural difference. For example, M. fuscus

    are less aggressive than R. fuscipes, they are slower, they use well-established runways and they nest

    communally above the ground during winter.

    Evidence of foxes was found in two of the 14 bogs surveyed, including Ginini West and Cotter Source.

    5.6 Conclusion

    M. fuscus is present at a range of sites in the higher elevation areas of the ACT where there is

    suitable habitat. Whilst the wildfire of 2003 burnt most of the habitat for the species, individuals

    survived in unburnt patches, and 13 years post-fire the species appears to still maintain its pre-fire

    distribution in the ACT. To what extent the 2003 fires reduced populations of M. fuscus in the ACT

    and resulted in the species going through a genetic ‘bottleneck’ is not known. Although it is not

    possible to determine a trend in abundance from a single ‘snapshot’ study, the high occupancy rate

    across the majority of bogs surveyed in the ACT subalpine region suggests that the populations of the

    species are currently relatively stable, albeit within a small and patchy distribution. On-going

    monitoring is required to determine whether ACT populations of M. fuscus are experiencing longer-

    term declines that appear to have occurred in other areas. This study provides baseline information

    on vegetation parameters and an index of density of M. fuscus and of introduced predators and

    competitors in bogs against which future changes can be assessed and key-threatening processes can

    be monitored. Population monitoring of M. fuscus and vegetation assessments undertaken every 5-

    10 years would enable long-term trends to be identified. The methods used in this study are an

    efficient and repeatable method for identifying trends in M. fuscus abundance and distribution.

    Acknowledgements

    We thank Melissa Snape, Luke Johnston and Sophia Callander for assistance in the field. We also

    thank Melita Milner, Felicity Grant, David Whitefield, Ken Green and Will Osborne for assistance.

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  • Distribution and ecology of the Broad-toothed Rat 25

    Appendix

    Figure A. Mastacomys fuscus runway.

  • 26 Distribution and ecology of the Broad-toothed Rat

    Figure B. Mastacomys fuscus scats.

    Figure C. Vegetation community dominated by medium heath.

  • Distribution and ecology of the Broad-toothed Rat 27

    Figure D. Vegetation community dominated by Poa.

    Figure E. Vegetation community dominated by sedge.

  • 28 Distribution and ecology of the Broad-toothed Rat

    Figure F. Woodland vegetation community.

    Figure G. Vegetation community dominated by low heath.